Ecology, 95(10), 2014, pp. 2860–2869
Ó 2014 by the Ecological Society of America
Movements vary according to dispersal stage, group size,
and rainfall: the case of the African lion
NICHOLAS B. ELLIOT,1,4 SAMUEL A. CUSHMAN,2 ANDREW J. LOVERIDGE,1 GODFREY MTARE,3
1
AND DAVID W. MACDONALD
1
Wildlife Conservation Research Unit, Department of Zoology, University of Oxford, Recanati-Kaplan Centre, Tubney House,
Abingdon Road, Tubney, Oxfordshire OX13 5QL United Kingdom
2
USDA Forest Service, Rocky Mountain Research Station, 2500 S Pine Knoll Drive, Flagstaff, Arizona 86001 USA
3
Zimbabwe Parks and Wildlife Management Authority, P.O. Box CY140, Causeway, Harare, Zimbabwe
Abstract. Dispersal is one of the most important life-history traits affecting species
persistence and evolution and is increasingly relevant for conservation biology as ecosystems
become more fragmented. However, movement during different dispersal stages has been
difficult to study and remains poorly understood. We analyzed movement metrics and patterns
of autocorrelation from GPS data for 20 lions (Panthera leo) over a five-year period. We
compared movement among different stages of natal dispersal (departure, transience, and
settlement), in addition to that of territorial adults of both sexes. The movement of lions
differed according to dispersal stage, sex, group size, and rainfall. As expected, during
dispersal lions moved faster and further and in a more directional manner than pre- or postdispersal. Transient movement was more directional than adult movement, but somewhat
surprisingly, was slower with less net displacement than that of territorial males. Interestingly,
the effect of group size on movement differed between transient males and territorial males;
solitary dispersers moved faster and further than individuals in bigger groups, while territorial
males had the opposite trend. Although our sample size is limited, our results suggest a
transition from directional movement during transience to random or periodic use of a fixed
territory after settlement. In addition, group size may affect the search and settlement
strategies of dispersers while seeking a territory in which to settle.
Key words: African lion; animal tracking; dispersal; Hwange National Park, Zimbabwe; movement;
Panthera leo; search strategies; seasonal variation; spatial autocorrelation.
INTRODUCTION
Dispersal is one of the most important life-history
traits involved in species persistence and evolution
(Clobert et al. 2001). As ecosystems become more
fragmented, dispersal is increasingly important as it is
often the only mechanism by which organisms can move
between populations and thus maintain genetically
diverse meta-populations (Clobert et al. 2012). Despite
its importance, little is known about dispersal movement
or the search strategies employed by dispersing individuals as they move through novel environments. This is
largely due to the practical difficulties associated with
data collection of this often once-in-a-lifetime event.
However, mortality rates, distances travelled, and
selection of settlement sites largely depend on the search
strategy of the disperser (Conradt et al. 2003, Doerr and
Doerr 2004, Conradt and Roper 2006). The paucity of
data on dispersal movement has resulted in most
population and evolution models assuming that individuals move at random in heterogeneous environments
Manuscript received 28 September 2013; revised 7 February
2014; accepted 26 February 2014; final version received 21
March 2014. Corresponding Editor: M. K. Oli.
4 E-mail: elliot.nb@gmail.com
(for a review see Patterson et al. 2008); however,
unrealistic representations of dispersal are likely to yield
inaccurate predictions regarding dispersal behavior and
processes (Bowler and Benton 2005). Furthermore,
recent evidence suggests that dispersal movements are
highly complex and may vary depending on the three
stages of dispersal (departure, transience, settlement;
Clobert et al. 2009, Delgado et al. 2009). A thorough
assessment of the ecological and evolutionary implications of dispersal thus requires robust empirical studies
that have been lacking and widely called for by recent
reviews (Jacobson and Peres-Neto 2010, Clobert et al.
2012, Baguette et al. 2013). Such studies will inform
conservation strategies (Macdonald and Rushton 2003)
and improve the way dispersal is represented in
simulation models (Zollner and Lima 2005) and
connectivity studies (Schwartz et al. 2009).
Shifting between dispersal stages brings about
changes in animal behavior (Gese 1998), particularly
in territorial species where territory holders and
dispersers show discernible ecological differences
(Campioni et al. 2012). In addition to dispersers being
relatively uninformed of key spatial and temporal
characteristics of the new environment, dispersers and
territory holders have different goals in that dispersers
2860
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MOVEMENT ECOLOGY OF A LARGE CARNIVORE
aim to establish a territory while avoiding conspecifics,
whereas established individuals need to maintain their
territory and defend access to mates (Campioni et al.
2010).
While research into movement patterns during dispersal is an emerging ecological field, the study of
movement ecology is vast and advanced (for reviews see
Holyoak et al. 2008, Schick et al. 2008) and can be
integrated into dispersal studies. For example, Schick et
al. (2008) argue that it is essential to understand the
interaction between environmental conditions and the
state of the organism to understand the drivers of
biologically based transition processes. Various empirical studies on vertebrates have highlighted behavioral
differences between the different dispersal phases in
birds (Stutchbury 1991, Delgado and Penteriani 2008),
reptiles (Aragón et al. 2006), and to a lesser extent,
mammalian carnivores (Gese 1998); however, there is a
lack of research linking these dispersal stages to
characteristics of the external environment, which is
imperative to understanding the drivers of movement
(Schick et al. 2008). To our knowledge, there has been
no research investigating seasonal movement patterns
during dispersal compared with territorial adults of the
same species and how movement varies depending on
group size. Doing so provides a unique opportunity to
evaluate the shifting search strategies of dispersers and
to obtain insight into the drivers of movement. To this
end, we studied the movement ecology of African lions
(Panthera leo) in all three dispersal phases, in addition to
territorial adults of both sexes.
Lions live in fission–fusion groups (Packer et al.
1990), and prides defend their territories. Adult males
do not tolerate the presence of non-coalition members,
with territorial encroachment usually resulting in
conflict (Grinnell et al. 1995, Packer 2001). Dispersal
in lions is sex biased, as subadult males always disperse,
while females are usually philopatric (Pusey and Packer
1987). During ‘‘departure,’’ most individuals conduct
prospecting searches outside their natal territory prior
to eventual departure (N. B. Elliot, unpublished data).
Little is known of the ‘‘transience’’ and ‘‘settlement’’
phases, but dispersing lions either settle in a vacant
area or challenge resident males for territory. While
many mammals, for example, the Eurasian badger
(Meles meles), disperse seasonally and over short
periods of time (Macdonald et al. 2008), lions may
disperse throughout the year, and the number of
transient months can be relatively prolonged, making
it possible to gather substantial quantities of data
during dispersal.
In this paper we investigate nine predictions (Table 1)
relating to the hypothesis that individual needs vary in
accordance with life stage, bringing about broad
behavioral changes, such as shifting movement patterns
and search strategies (Zollner and Lima 1999, Van Dyck
and Baguette 2005, Zollner and Lima 2005, Stamps
2006, Schick et al. 2008, Penteriani et al. 2011). To
2861
TABLE 1. Predictions of hypotheses relating to the movement
of African lions (Panthera leo) in different demographic
categories and how their movement is influenced by rainfall
and group size.
Data set and predictions
Comparing subadult dispersal stages
1) Transient lions move fastest and furthest
2) Transient lions display most directional
movement
3) Post-dispersal lions cease directional movements
and display random and periodic movement
Source
1, 2
3, 4, 5
7
Comparing transients and territorial adults
4) Transients move slowest and less far per night
6
5) Transient individuals display more directional
3
movement
6) Territorial adults predominantly display
7
periodic movements
7) Territorial adults respond to rainfall while
8, 9, 10
dispersers do not
8) Transients in smaller groups move more per
11, 12
night compared to larger groups
9) Territorial adults in bigger groups move further 8
per night compared to smaller groups
Notes: We analyzed two different data sets: The first
consisted of pre-dispersal, transient, and post-dispersal lions;
the second consisted of transients, territorial males, and
territorial females. Type in boldface represents supported
predictions, and type in italics indicates partially or weakly
supported predictions. Predictions were based on hypotheses
and findings from the following sources: 1, Delgado et al. 2009;
2, Campioni et al. 2012; 3, Zollner and Lima 1999; 4, Van Dyck
and Baguette 2005; 5, Stamps 2006; 6, Zollner and Lima 2005;
7, Valeix et al. 2010; 8, Loveridge et al. 2009; 9, Campioni et al.
2010; 10, Vuilleumier and Perrin 2006; 11, Packer et al. 1988;
and 12, Baguette and Van Dyck 2007.
achieve this, we used an extensive data set based on five
years of fine-scale GPS data collected on 20 lions of
varying status. We analyzed patterns of autocorrelation
and classified movement as directional, periodic, or
random, in addition to calculating various movement
metrics. We then examine whether movement of lions
differs depending on their demographic category (predispersal, transient, post-dispersal, territorial adult), sex,
group size, and rainfall.
METHODS
Study area
The study area (;7000 km2) was located in the
northern section of Hwange National Park (HNP),
Zimbabwe (19800 0 S, 26830 0 E). HNP covers ;15 000
km2 of semiarid savannah. Vegetation consists primarily
of woodland and bushland savannah (64%), and
communities are dominated by Baikiaea plurijuga,
Colophospermum mopane, Combretum spp., Acacia
spp., and Terminalia sericea (Rogers 1994). The longterm mean annual rainfall of 613 mm is highly variable
(CV ’ 26%) and generally falls between October and
April. Surface water is available from seasonal waterholes, although only a few hold water in the dry season,
during which time water is artificially supplied to some
(;50) waterholes.
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NICHOLAS B. ELLIOT ET AL.
Ecology, Vol. 95, No. 10
FIG. 1. Example Mantel correlograms that were visually classified by assessing the first half of the correlogram. These plots
illustrate an individual that shifted from periodic to directional to random movement in three consecutive windows. See Methods
and Appendix A for details on correlogram classification.
Lion movement data
Between 2007 and 2012, we obtained movement data
from 20 lions in different social groups. Each was fitted
with a GPS enabled radio-collar (see Loveridge et al.
2007 for details) pre-programmed to take hourly fixes
when lions are active (18:00–07:00 hours).
Data preparation
To enhance accuracy, only fixes with a dilution of
precision (DOP) , 10 were retained for analysis (Frair
et al. 2010). This resulted in a data set of 40 669 locations
for subadult males (93% DOP , 5), 72 030 locations for
territorial males (96% DOP , 5), and 60 139 locations
for territorial females (97% DOP , 5).
To reduce the effects of non-stationarity (Cushman et
al. 2005), the study period was split into sequential,
nonoverlapping temporal windows of 30-day intervals.
In total we created 56 window periods, the first starting
1 October 2007 and the last on 22 May 2012. The data
for each individual lion were split according to these
windows, resulting in a total data set of 396 windows.
Mantel correlograms
Mantel correlograms reveal patterns of spatial and
temporal autocorrelation that are highly informative in
elucidating scales and patterns of ecological processes
(Legendre 1993) and provide a detailed picture of
movement variability (Cushman et al. 2005). To identify
different movement patterns for each window and
individual, we calculated two matrices (distance and
time): The distance matrix was produced by calculating
geographical distances between each pair of locations,
and the time matrix was computed by calculating the
difference in decimal days between each pair of
locations. We used Mantel tests to assess the level of
association between these two matrices (Mantel 1967).
We computed Mantel correlograms (Oden and Sokal
1986) to assess the levels of spatial autocorrelation in
lion movements across a range of lag times. We visually
classified the correlogram ‘‘shape’’ for each window (for
examples, see Fig. 1). Cushman (2010) simulated three
path types under 18 different movement rules and
showed that each path type had a characteristic
autocorrelation structure and correlogram shape. Thus,
correlogram shapes are diagnostic of broad movement
patterns and provide an accurate and repeatable method
to classify paths into categories such as directional,
random, and periodic movement (Cushman et al. 2005,
Cushman 2010). In the current study, we used the shape
of the first half of each correlogram to classify the
movement during that period into three ‘‘shape’’
categories: directional, periodic, or random movement.
Directional movement, akin to a correlated random
walk, is typified by a constant cline in the correlogram
from positive to negative autocorrelation at increasing
time lags, indicating a pattern of movement in which
successive locations in time become farther apart in
space. Periodic movement, resulting from central place
random walks or periodic revisits to a collection of
locations, is characterized by repeated cycles between
strong positive and strong negative autocorrelation.
Random movement within a fixed home range, akin to
bounded correlated random walks, is typified by a rapid
drop from positive to negative autocorrelation and
subsequent fluctuation near zero autocorrelation (see
Fig. 1 for sample correlograms; for full explanation of
methods, see Appendix A, and Cushman et al. 2005,
2010).
Movement metrics
We calculated three movement metrics: speed, path
length, and net displacement. Speed (m/h) was calculated by dividing the total distance travelled per night
by the number of hours. Path length was the total
October 2014
MOVEMENT ECOLOGY OF A LARGE CARNIVORE
distance travelled each night, and net displacement was
the distance between the first and last fix on a given
night. Each parameter was then averaged per 30-day
window.
2863
by deaths or increased by the addition of new members
into a coalition or pride. For adult group size, we only
included individuals over 36 months of age, while
transient groups could be younger.
Rainfall
Statistical analyses
Daily rainfall was recorded at two weather stations
within the study area, approximately 30 km apart. The
figures were averaged to give a rainfall profile across the
study area. We did not expect that rainfall in itself would
impact lion movements, but rather, that the replenishment of water holes, vegetation growth, and subsequent
prey dispersion would influence lion movement patterns.
We therefore computed total rainfall for 60 days prior to
the start of each window.
The structure of our data necessitated different
modeling approaches for Mantel correlogram shape
(categorical response) and movement metrics (continuous response). Furthermore, we analyzed two different
data sets that consisted of: (1) the subadult dispersal
stages (pre-dispersal, transient, and post-dispersal); and
(2) the transients, territorial males, and territorial
females.
Demographic categories
To assess the relationships between correlogram shape
and demographic parameters and rainfall, we used
generalized linear mixed models using combinations of
two shape types per analysis (i.e., three sets of pairs). An
example analysis consists of a response variable (e.g.,
periodic [0] or directional [1] movement), each of the
fixed effects (demographic category, group size, and
rainfall), their interaction terms and lion identity as a
random intercept (see Appendix B for models and model
selection statistics). These analyses were conducted in R
2.15.1 (R Development Core Team 2012), package lme4
v.0.9-0 (Bates et al. 2012) using a binomial error
structure and logit-link function, with estimates provided on the logit scale.
To analyze the movement metrics, we fit mixed-effects
linear models with a continuous response variable
(either speed, path length, or net displacement), each
of the fixed effects (demographic category, group size,
and rainfall), their interaction terms and lion identity as
a random intercept (see Appendix C for models and
model selection statistics). We used R package nlme
v.3.1 (Pinheiro et al. 2012).
The data used in this study were from lions in three
demographic categories: territorial males (n ¼ 6),
territorial females (n ¼ 5), and subadult males (n ¼ 9).
To assess changes in movement during dispersal, the
subadult data were further split into the three phases of
dispersal (Clobert et al. 2001): pre-dispersal (n ¼ 4),
transient (n ¼ 9), and post-dispersal (n ¼ 5). Based on
field observations, we classified the demographic categories as follows.
Pre-dispersal.—Four subadult males were collared
while still with their natal pride. We were interested in
investigating movements leading up to final departure
and so only used data six months prior to dispersal (18
windows).
Transience.—The timing of dispersal was calculated as
the midpoint between the last time a subadult was seen
with its natal pride and the first time either was seen
alone (mean time interval ¼ one month; SD 6 1.52).
Nine individuals made up the transient data set (58
windows).
Post-dispersal.—Establishment of a territory was
deemed to have occurred when a transient subadult
had been in a consistent home range for a minimum
period of two months and continued to have a fixed
home range thereafter. The onset of post-dispersal was
then back-dated to the time at which the lion entered the
new home range. We were interested in movements
shortly after establishment and therefore discarded all
data that extended beyond six months after settlement.
Five transient males were deemed to establish territories
(27 windows).
Territorial adults.—Lions in the study area have been
closely monitored since 2002. All adults in this study
were known to be territorial through direct observation
and GPS data. In total we used 158 windows for
territorial males and 135 for territorial females.
Group size
All lions were closely monitored throughout the study
period allowing for accurate recording of group size.
Group size varied over the study period, being reduced
Movement analysis
Data sets
First we compared the subadult data set during the
three dispersal phases. Our primary interest in this
data set was to assess change in movement depending
on demographic category irrespective of group size or
rainfall. Furthermore, our data set of pre- and postdispersal was relatively small, necessitating simple
models. For each movement metric and type, we
constructed models with demographic category as the
only explanatory variable (predictions 1–3 in Table
1).
Second we analyzed the data set comprising territorial
males, females, and transient males (without the
pre- and post-dispersal data). We assessed the relationship between movement and demographic category,
group size, and rainfall. To investigate predictions 4–9
(Table 1), we created 12 a priori candidate models
including each of the main effects and their interaction
terms (Appendices B and C). The most complex model
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NICHOLAS B. ELLIOT ET AL.
TABLE 2. Estimated parameters and 95% confidence intervals
(CI) from mixed models assessing variation in movement
between transient, and pre- and post-dispersal lions in
Hwange National Park, Zimbabwe.
Response variable
and parameter
95% CI
Estimate
SE
Lower
Upper
Net displacement
Transient
Pre-dispersal
Post-dispersal
4336.2
1225.1
743.2
304.6
460.2
392.9
3740.2
2125.7
1512.2
4932.3
324.4
25.8
Path length
Transient
Pre-dispersal
Post-dispersal
5942.6
1239.8
298.6
490.4
586.8
498.2
4982.9
2388.2
1273.5
6902.3
91.4
676.3
Speed
Transient
Pre-dispersal
Post-dispersal
547.7
106.0
47.4
33.6
45.8
39.0
481.8
195.7
123.7
613.5
16.3
29.0
Directional vs. random
Transient
Pre-dispersal
Post-dispersal
0.8
1.6
2.7
0.3
0.8
0.8
0.1
3.1
4.4
1.5
0.1
1.1
Directional vs. periodic
Transient
Pre-dispersal
Post-dispersal
0.3
1.2
2.0
0.3
0.7
0.8
0.3
2.7
3.6
0.8
0.2
0.4
Random vs. periodic
Transient
Pre-dispersal
Post-dispersal
0.5
0.8
0.4
0.4
0.5
0.6
0.2
1.8
1.6
1.2
0.3
0.9
considered was: demographic 3 group size þ rainfall 3
demographic. We used the same 12 a priori candidate
models to analyze all three movement metrics and
correlogram shapes. Model selection was used to
identify the best model based on Akaike information
criterion corrected for small sample size (AICc). We
decided a priori that if one model was clearly superior
(wi . 0.9), this would be used, otherwise we would
Ecology, Vol. 95, No. 10
average parameter estimates across models with AICc
differences (Di , 3), correcting for model weights using
R package AICcmodavg v.1.30 (Burnham and Anderson 2002, Mazerolle 2013). R syntax for all analyses and
for computation of Mantel correlograms are provided in
the Supplement.
RESULTS
Subadult male dispersal stages
Transient lions had higher net displacement and
moved further and faster per night than either pre- or
post-dispersal (prediction 1 in Table 1). This difference
was largest between transient and pre-dispersal lions
(Table 2, Fig. 2). Post-dispersal lions had the second
highest values of these parameters, while pre-dispersers
had the lowest.
Transient lions were more likely to use directional
rather than random or periodic movement compared to
pre- and post-dispersal lions (predictions 2–3 in Table 1;
see Table 2 and Fig. 3 for comparisons).
Transients and territorial adults
Lion movement varied depending on demographic
category, rainfall, group size, or a combination of these.
There was no top model (wi , 0.9) in our analyses of
movement metrics (speed, path length, net displacement)
or correlogram shape; thus, all reported parameter
estimates were obtained by averaging across models
with AICc differences ,3 from the top model, correcting
for model weights (see Appendices B and C for model
selection statistics and model averaging procedure). The
most supported candidate models for all movement
metrics consisted of the interaction terms group size 3
demographic and rainfall 3 demographic, in addition to
their main effects. The most supported candidate models
in our analysis of correlogram shape contained only the
main effects (demographic, rainfall, and group size).
FIG. 2. Differences in movement patterns between pre-dispersal, transient, and post-dispersal African lions (Panthera leo) in
Hwange National Park, Zimbabwe, in relation to (a) net displacement, (b) path length, and (c) speed. Boxes show medians, 25%,
and 75% quartiles. Triangles indicate means. Whiskers indicate 10th and 90th percentiles. Dots represent the raw data.
October 2014
MOVEMENT ECOLOGY OF A LARGE CARNIVORE
2865
(estimate ¼ 2307 6 634, 95% CI ¼ 976–3637) per night,
while females moved slower (estimate ¼133 6 52, 95%
CI ¼ 242 to 24). Similarly, territorial males had
higher net displacement (estimate ¼ 880 6 343, 95% CI ¼
160–1600), while females had lower net displacement
(estimate ¼ 1569 6 356, 95% CI ¼ 2317 to 821)
compared to transient subadult males (prediction 4 in
Table 1).
Transient males were more likely to use directional
rather than periodic movement (Fig. 4) compared to
territorial males (estimate ¼ 1.3 6 0.5, 95% CI ¼ 2.3
to 0.3). Transient males were also more likely to use
directional movement rather than random movement
compared to territorial males (estimate ¼1.9 6 0.6, CI
¼ 3 to 0.8) and territorial females (estimate ¼ 1.7 6
0.6, CI ¼ 2.8 to 0.5; predictions 5 and 6 in Table 1).
Although group size and rainfall were included in the
top models (Appendix C), there was no clear trend
(estimate and 95% CI included 0).
Rainfall
FIG. 3. Differences in correlogram shape in relation to predispersal, transient, and post-dispersal lions in Hwange
National Park. Correlograms were classified as long-range
directional movement (directional), cyclical visits to focal areas
(periodic), or random use within a home range (random). The
width of segments is proportional to the amount of data for
each dispersal stage. See Fig. 1 and Appendix A for further
clarification.
Demographic categories
Compared to transient subadult males, territorial
males moved faster (estimate 6 SE [all estimates shown
with 6SE] ¼ 164 6 49, 95% CI ¼ 60–268) and further
The way in which lions responded to rainfall
depended on an interaction with demographic category.
All demographic groups moved slower, less far, and with
a lower net displacement with increasing rainfall, with
the effects generally largest for territorial females
(prediction 7 in Table 1; Table 3).
Group size
Group size ranged from one to four in adults and one
to three in transient males. Movement metrics for
territorial males, territorial females, and transient males
differed depending on group size (Table 4, Fig. 5).
Transient males in smaller groups moved faster, and had
higher net displacement and higher path length than
those in bigger groups (prediction 8 in Table 1). In
contrast, territorial males displayed the opposite trend:
Bigger groups moved faster, and had higher net
displacement and a higher path length than smaller
TABLE 3. Model-averaged slope estimates from results of
mixed-effects linear models investigating lion movement
metrics (net displacement, path length, and speed) in relation
to rainfall in Hwange National Park.
FIG. 4. Differences in correlogram shape according to lion
demographics in Hwange National Park. Correlograms were
classified as long-range directional movement (directional),
cyclical visits to focal areas (periodic) or random use within a
home range (random). The width of segments is proportional to
the amount of data for each demographic. See Fig. 1 and
Appendix A for further clarification.
95% CI
Response variable
and interaction
Slope
SE
Lower
Upper
Net displacement
Rain 3 transient males
Rain 3 adult males
Rain 3 adult females
0.28
0.09
1.51
1.12
0.86
0.98
2.47
1.78
3.44
1.90
1.61
0.41
Path length
Rain 3 transient males
Rain 3 adult males
Rain 3 adult females
1.33
1.21
2.92
1.53
1.15
1.33
4.33
3.47
5.52
1.68
1.05
0.32
Speed
Rain 3 transient males
Rain 3 adult males
Rain 3 adult females
0.15
0.17
0.21
0.10
0.07
0.08
0.35
0.30
0.36
0.04
0.03
0.05
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NICHOLAS B. ELLIOT ET AL.
TABLE 4. Model-averaged slope estimates from results of
mixed-effects linear models investigating lion movement
metrics (net displacement, path length, and speed) in relation
to group size in Hwange National Park.
95% CI
Response variable
and interaction
Slope
SE
Lower Upper
Net displacement
Group size 3 transient males 1494 320 2121
Group size 3 adult males
405 121
168
Group size 3 adult females
394 156 701
867
642
88
Path length
Group size 3 transient males 2085 482 3029 1140
Group size 3 adult males
674 171
339
1009
Group size 3 adult females
130 226 573
313
Speed
Group size 3 transient males
Group size 3 adult males
Group size 3 adult females
144
37
11
39
13
18
220
11
46
68
63
23
groups. Territorial females exhibited the same tendency
as transient males, but increasing group size had a less
marked effect as there was only substantial decrease in
net displacement (prediction 9 in Table 1; Table 4).
DISCUSSION
Our findings show the extent to which lion movement
is influenced by group size and rainfall and how this
differs depending on demographic category. By incorporating an environmental variable in addition to the
dispersal stages of the organism (Schick et al. 2008), we
offer a unique insight into differential responses to
shifting environmental conditions and group size depending on the life stage and sex of a species.
Transient lions exhibited directional movement, while
adult movement was primarily a composite of random
and periodic movements. Differences in movement are,
to a certain extent, determined by an individual’s ability
Ecology, Vol. 95, No. 10
to perceive its surroundings and acquire knowledge
(Vuilleumier and Perrin 2006). As such, dispersers
moving through novel environments may move differently relative to territorial individuals that have learned
to maximize the resources within that patch, as was
reported with transient eagle owls (Bubo bubo; Delgado
et al. 2009). The observed periodic movement in adults
was expected, given that, in this ecosystem, waterholes
are a key locus for lions; prey select areas close to
waterholes (Valeix et al. 2009) and subsequently so do
lions (Valeix et al. 2010), which appear to periodically
rotate between waterholes (prediction 6 in Table 1;
Valeix et al. 2011).
Pre-dispersal lions exhibited periodic and random
movement patterns shifting to high levels of directional
movement during dispersal (prediction 2 in Table 1).
This accords with Zollner and Lima’s (1999) simulated
dispersal experiments and subsequent predictions: They
found that straighter paths vastly improved the probability of survival and that this type of movement was the
most effective search rule as straighter paths avoid
redundant search and improve the likelihood of finding
a vacant patch quickly (prediction 5 in Table 1). With
territorial species such as lions, it is plausible that, unless
a disperser is going to challenge for a territory, he will
pass through in a directional manner, and minimize the
chance of an aggressive encounter with the resident
adult. Such directional movement suggests a sequential
search strategy whereby individuals either accept or
reject patches they encounter; if rejected, they continue
the search in a directional manner, and do not revisit
patches (comparative search strategies; Stamps et al.
2005, Stamps 2006). Consistent with this idea, the
amount of directional movement declined dramatically
once establishment occurred, replaced by random and
periodic movements (prediction 3 in Table 1). In
contrast, territorial adults displayed higher proportions
FIG. 5. Relationship between demographic category and group size in relation to net displacement of lions in Hwange National
Park. There were no groups of four in transient lions. Boxes show medians, 25%, and 75% quartiles. Triangles indicate means.
Whiskers indicate 10th and 90th percentiles. Dots represent the raw data.
October 2014
MOVEMENT ECOLOGY OF A LARGE CARNIVORE
of directional movement than did post-dispersal lions.
We speculate that during establishment, a subadult lion
consolidates a patch, around a waterhole for instance,
leading to exaggerated patterns of periodic and bounded
home range movements, and as it becomes familiar with
and secure in its new territory, it switches to movement
patterns similar to those of territorial adult males.
Additional data on post-dispersal movements may
confirm this.
Our findings concur with what Van Dyck and
Baguette (2005) termed ‘‘special movements’’ in relation to directional dispersal movements designed for
displacement ( prediction 5 in Table 1). However,
‘‘special movements’’ are associated with high speed
as reported for transient eagle owls that moved faster
and straighter than established individuals (Delgado et
al. 2009). In contrast, our results showed that transient
males were slower and moved less than territorial males
(prediction 4 in Table 1). One explanation is that by
using directional movement at a decreased locomotory
rate, transient males effectively search the area while
limiting their detection probability. Zollner and Lima
(2005) demonstrated that transient individuals can
benefit from slower speeds due to more effective antipredator behavior. More tortuous paths covering a
larger area would entail leaving more scent, which
would increase the likelihood of detection by territorial
males. Indeed, increased activity has been shown to
increase predation susceptibility (Ebenhard 1987) and
predation rates (Norrdahl and Korpimäki 1998) in
small mammals.
Depending on their demographic, group size influenced how individuals moved. When in small groups,
transient lions moved faster and farther (prediction 8 in
Table 1), as did territorial females, although the effect
was less marked. In contrast, territorial males had the
opposite trend (prediction 9 in Table 1). Fluctuations
in lion home range size are strongly suggestive of
expansionism (Kruuk and Macdonald 1985), as home
range size increases with increasing pride biomass
(Loveridge et al. 2009). Thus, with larger territories
to defend, males in bigger groups have increased
movement compared to smaller groups in smaller
territories. On the other hand, females are not as active
as males in patrolling their territories (Schaller 1972),
and thus, while bigger groups of females have larger
territories, their movements may reflect a strategy of
food acquisition (which is concentrated around waterholes) and rearing of offspring. Lions in larger
coalitions are more likely to gain residence in a pride
(Bygott et al. 1979, Packer et al. 1988), and among
transients it is plausible that a solitary disperser has
little chance of displacing the resident males and would
benefit from moving directly through an occupied
territory. Thus, the decreased locomotion of transient
individuals in bigger groups could reflect ‘‘boldness’’ as
opposed to ‘‘shyness’’ (Baguette and Van Dyck 2007).
While it has been reported that bold individuals
2867
dispersed farther (Fraser et al. 2001) with larger and
straighter movements (Delgado and Penteriani 2008),
in our scenario the opposite could apply. Bigger groups
may face fewer consequences if detected by resident
males and can afford to remain within occupied
territories for longer periods, thereby challenging for
territory. Solitary individuals, on the other hand,
represent the ‘‘shy’’ individuals and are frequently
displaced from occupied territories causing them to
move faster, with higher net displacement per night
through the landscape while searching for a (often rare)
vacant territory, and may be less inclined to challenge
for one.
Lions showed differing response to rainfall in the
preceding 60 days depending on their demographic
(prediction 7 in Table 1). While the changing of seasons
has no effect on lion home range size of either sex in
HNP (Loveridge et al. 2009), we show that when using a
more fluid approach to climatic data (i.e., rainfall as
opposed to pre-defined seasons delineated by calendar
months), variation in movement is detected. The slopes
defining these relationships are shallow (Table 4), and
our small sample size necessitates caution; however,
these findings confirm our expectations. For instance,
that females decreased their speed and path length with
increased rainfall may be explained by the filling of
waterholes and the addition of small cubs, which may
constrain the movements of females. Cubs were more
common in the wet season during this study. Indeed,
cubs under six months of age were present in 49% of all
female wet-season windows compared to 38% of dryseason windows.
In summary, our findings reveal that in territorial
species such as lions, movement may be influenced by
the sex and dispersal stage of an individual. This
variation may be further explained by incorporating
group size and environmental variables. Our results
indicate a transition from directional exploratory
strategies during dispersal to restricted random and
periodic movement during settlement, followed by more
expansive movement after establishment. As ecosystems
become increasingly fragmented, it is crucial to elucidate
the patterns and processes of movement during the
different stages of dispersal to better understand the
associated shifting behaviors. This will improve our
understanding of how species may persist in fragmented
landscapes and how fencing, for example, may curtail
and alter the movements described in this paper. Our
results provide the first investigation of seasonal
movement patterns during dispersal in conjunction with
territorial adults and how this varies depending on
group size. There has been wide recognition of the
importance of empirical studies on dispersal movements,
and we hope that our results will improve the way
dispersal is represented in simulation models (e.g.,
Zollner and Lima 2005) and connectivity studies (e.g.,
Schwartz et al. 2009).
2868
NICHOLAS B. ELLIOT ET AL.
ACKNOWLEDGMENTS
The Director General of the Zimbabwe Parks and Wildlife
Management Authority is acknowledged for providing the
opportunity to carry out this research and for permission to
publish this article. We are grateful for the assistance of B.
Stapelkamp, J. Hunt, and the ecological staff of HNP in
undertaking fieldwork for this study. We thank S. Périquet for
useful comments and F. Broekhuis for guidance throughout the
preparation of the manuscript. We also thank J. Fryxell and
other reviewers for helping improve the manuscript. HNP Lion
Project was supported by the Darwin Initiative for Biodiversity
Grant, the very kind generosity of J. Cummings and family, and
A. Gardiner and the following Foundations: Eppley, Lillian
Jean Kaplan, RG Frankenberg, Rufford, Boesak-Kruger, and
Disney.
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SUPPLEMENTAL MATERIAL
Appendix A
A detailed description for classifying correlogram ‘‘shape’’ into directional, periodic, or random movements, also including
background information on the method and a simulation to further illustrate it (Ecological Archives E095-247-A1).
Appendix B
Model selection statistics for generalized linear mixed models investigating correlogram shape (directional, periodic, and random
movements) of lion movement paths in Hwange National Park (Ecological Archives E095-247-A2).
Appendix C
Model selection statistics for mixed-effects linear models investigating lion movement parameters (net displacement, path length,
and speed) in Hwange National Park (Ecological Archives E095-247-A3).
Supplement
Sample R code of all analyses in the manuscript in addition to R code and instructions for computing mantel correlograms
(Ecological Archives E095-247-S1).